J OURNAL OF C RUSTACEAN B IOLOGY, 32(3), 457-464, 2012
DISTRIBUTION PATTERNS OF THE AMERICAN SPECIES OF THE FRESHWATER
GENUS EUCYCLOPS (COPEPODA: CYCLOPOIDA)
Nancy F. Mercado-Salas 1,∗ , Carmen Pozo 1 , Juan J. Morrone 2 , and Eduardo Suárez-Morales 1
1 El
Colegio de la Frontera Sur (ECOSUR), Unidad Chetumal, Av. Centenario km 5.5, Chetumal,
77014 Quintana Roo, Mexico
2 Museo de Zoología “Alfonso L. Herrera”, Departamento de Biología Evolutiva, Facultad de Ciencias,
Universidad Nacional Autónoma de México (UNAM), Apdo. Postal 70-399, 04510 Mexico, D.F., Mexico
ABSTRACT
Based on the superposition of 19 individual tracks of American species of the freshwater copepod genus Eucyclops, two generalized
tracks were found. The Western Amazonian track (southern Peru, eastern Brazil, and central Colombia) corresponding to the Amazonian
subregion and the South American Transition Zone, and the Mesoamerican-Northwestern South American track (central Colombia, Central
America, and northeastern Mexico) corresponding to the Neotropical region, the Mexican Transition Zone, and the Nearctic region.
One node was found in Colombia, an area where both generalized tracks intersect. The distributional patterns of Eucyclops apparently
involve two cenocrons: one Holarctic, and another Paleotropical. The Western Amazonian generalized track can be correlated with the
existence of rivers that function either as barriers or dispersal passageways, the uplift of the Andes, and the presence of the Miocene
“Pebas lake/wetland system.” The Mesoamerican-Northwestern South American generalized track can be associated with climate changes
resulting from the uplift of North American mountain ranges, the presence of marine barriers (Isthmus of Tehuantepec and Panama) and
the uplift of mountains in southern Mexico and Central America. The closing of the marine barrier represented by the Isthmus of Panama
seems to have been a key event in the northward and southward dispersal of Eucyclops in the Americas.
K EY W ORDS: copepods, Cyclopinae, Eucyclops, generalized tracks, individual tracks, panbiogeography
DOI: 10.1163/193724012X626502
I NTRODUCTION
The freshwater copepod genus Eucyclops Claus, 1893 comprises 106 nominal species and subspecies (Dussart and Defaye, 2006). It is one of the most taxonomically challenging
genera in Cyclopidae, containing several problematic taxa
and some species groups with a high intraspecific variability. This, together with incomplete descriptions, has generated a taxonomic history that includes many species with
an uncertain status (Collado et al., 1984; Reid, 1985; Ishida,
1997) and a complex taxonomy that relies on only a few relatively stable characters. The species of Eucyclops are divided
into three subgenera: Eucyclops sensu stricto, which contains most of the known species; Stygocyclops Pleša, 1971,
with a single species (E. [S.] teras [Graeter, 1907]) from
Switzerland, and Isocyclops Kiefer, 1957, which includes
two species endemic to Lake Tanganyika (Dussart and Defaye, 2001, 2006; Suárez-Morales, 2004). In the Americas,
there are more than 800 records of the genus, corresponding to 28 nominal species, most of which are distributed in
eastern United States, Mexico, Argentina, and Brazil.
Few works have analyzed the biogeographic affinities of
the freshwater copepods of the New World. Menu-Marque
et al. (2000) studied the species of Boeckella Guerne and
Richard, 1889 using a track analysis. They found that their
biogeographical patterns reflect the existence of an ancient
Austral biota, the biotic evolution of which was influenced
∗ Corresponding
greatly by the break-up of the Gondwana supercontinent,
this genus is also distributed in Australia and New Zealand.
Suárez-Morales et al. (2004) analyzed the distributional
patterns of the cyclopoid species of the Yucatan Peninsula
and suggested that it reflects post-Pliocene events, with
a major Neotropical biotic influence. In a phylogenetic
analysis of all known species of Mesocyclops Sars, 1914,
Hołyńska (2006) highlighted the high level of endemism
of this genus in South America because of its isolation
during the Cretaceous, which allowed the preservation of
ancient lineages. De los Ríos et al. (2010) analyzed the
distributional patterns of the Chilean cyclopoids, finding
some species endemic to the Atacama and Magellanic
Moorland biogeographic provinces, and others reported in
several areas in South America.
Evolutionary biogeography integrates distributional, phylogenetic, molecular, and paleontological data in order to
discover biogeographic patterns and assess the historical
changes that shaped them (Morrone, 2009). It follows a series of steps that include: 1) identification of biotic components, which are sets of spatio-temporally integrated taxa
that coexist in given areas, graphically represented as generalized tracks and areas of endemism; 2) testing relationships
between biotic components, with help of cladistic biogeography, which uses information on the cladistic relationships
between taxa and their geographic distribution to postulate
author; e-mail: [email protected]
© The Crustacean Society, 2012. Published by Koninklijke Brill NV, Leiden
DOI:10.1163/193724012X626502
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 3, 2012
hypotheses on relationships between areas; 3) regionalization, which implies the recognition of successively nested
areas that allow a biogeographic classification; 4) identification of cenocrons, which are sets of taxa that share the
same biogeographic history, constituting identifiable subsets
within a biotic component by their common biotic origin and
evolutionary history; and 5) construction of a biotic scenario,
by accounting biological and non-biological data, which are
used to integrate a plausible scenario to help explain the
episodes of vicariance or biotic divergence and dispersal or
biotic convergence (Morrone, 2009).
The recognition of biotic components, the first step of an
evolutionary biogeographic analysis, can be done using the
panbiogeographic approach, which emphasizes the spatial
or geographic dimension of biodiversity to allow a better
understanding of evolutionary processes. Morrone (2006),
after a series of panbiogeographic analyses, recognized 70
biotic components in Latin America and the Caribbean region that were deemed as biogeographic provinces. These, in
turn, were grouped into major biotic components considered
as regions, subregions, and dominions; also, two transition
zones were distinguished. These biogeographic provinces
have been recognized by different authors analyzing several
groups including mammals, tropical snakes, fish parasites,
freshwater decapods and copepods, and terrestrial arthropods (Lopretto and Morrone, 1998; Menu-Marque et al.,
2000; Márquez and Morrone, 2003; Abrahamovich et al.,
2004; Escalante et al., 2004; Morrone and Márquez, 2008;
Yáñez-Ordoñez et al., 2008; Rosas-Valdéz and Pérez-Ponce
de León, 2008; Arzamendia and Giraudo, 2009; Asian et al.,
2010; Maya-Martínez et al. 2011).
Herein we analyze the geographical distribution of the
species of Eucyclops in the Americas using a track analysis,
which has been previously applied to other crustacean taxa
(Morrone and Lopretto, 1994; Lopretto and Morrone, 1998;
Menu-Marque et al., 2000). We identify individual and
generalized tracks, in order to contribute to the knowledge
of the spatial evolution of this copepod taxon.
M ATERIALS AND M ETHODS
Data
Distributional data were obtained from the literature (Marsh,
1893; Juday, 1915; Kiefer, 1926, 1929, 1931, 1934, 1936,
1956; Pearse and Wilson, 1938; Osorio-Tafall, 1943;
Comita, 1950; Lindberg, 1955; Robertson and Gannon,
1981; Collado et al., 1984; Dussart, 1984; Reid, 1985;
Suárez-Morales et al., 1985; Dussart and Frutos, 1986; Defaye and Dussart, 1988; Zamudio-Valdez, 1991; Reid and
Marten, 1995; Zannata-Juárez, 1995; Dodson and SilvaBriano, 1996; Santos, 1997; Grimaldo-Ortega et al., 1998;
Suárez-Morales and Reid, 1998; Gutiérrez-Aguirre, 1999;
Álvarez-Silva and Gómez-Aguirre, 2000; Elías-Gutiérrez,
2000; Fiers et al., 2000; Reeves, 2000; Barbiero et al.,
2001; Ishida, 2001; Rodríguez-Almaráz, 2002; Carling et
al., 2004; Suárez-Morales, 2004; Bruno et al., 2005; Frisch
et al., 2005; Alekseev et al., 2006; Gaviria and Aranguren,
2007; Elías-Gutiérrez et al., 2008; Jiménez-Trejo and
Vásquez-Vargas, 2008; Mercado-Salas, 2009; SuárezMorales and Walsh, 2009; De los Ríos et al., 2010). Additional records were obtained from the collection databases
of the Smithsonian Institution-National Museum of Natural
History and the zooplankton collection of El Colegio de la
Frontera Sur-Unidad Chetumal, ECO-CH-Z.
For some records lacking geographical data, localities
were geo-referenced with the aid of Google-Earth. Other
species, including Eucyclops macrurus (Sars, 1863), E.
neotropicus Kiefer, 1936, E. ariguanabaensis Brehm, 1948,
E. chilensis Löffler, 1961 (Menu-Marque and Locascio,
2011), E. siolii Herbst, 1962, E. breviramatus Löffler, 1963,
E. neomacruroides Dussart and Fernando, 1990, E. borealis
Ishida, 2001, and E. cuatrocienegas Suárez-Morales and
Walsh, 2009 have been cited for a single locality each.
These records do not provide relevant information for the
track analysis and were thus excluded. The records of E.
pectinifer (Cragin, 1883), E. serrulatus (Fischer, 1851), and
E. agilis (Koch, 1838) were a priori treated as referring to a
single species, following Alekseev et al. (2006). Eucyclops
agilis has been considered as a valid species by several
authors in the Americas, but its original description is
not accurate enough to consider it as a valid species; it
is currently recognized as a synonym of E. serrulatus
(Alekseev et al., 2006). Eucyclops serrulatus is distributed in
North Africa, the Mediterranean region, Europe, Russia, and
probably also in Central Asia. Records of the species in the
Americas should be carefully reexamined before assigning
them to the American form E. pectinifer, because some
of them could belong in fact to E. serrulatus. According
to Alekseev (personal communication to NFM-S, 2011)
the latter may have been introduced in the Americas by
human agency, other records could be E. pectinifer or even
undescribed species. Records of E. speratus (Lilljeborg,
1901), E. elegans (Herrick, 1884), and E. solitarius (Herbst,
1959) were merged into a single species, following Reid and
Marten (1995). After this process we obtained a database
of 446 records for E. pectinifer, E. delachauxi (Kiefer,
1925), E. silvestri (Brian, 1927), E. neumani neumani
(Pesta, 1927), E. prionophorus Kiefer, 1931, E. bondi
Kiefer, 1934, E. ensifer Kiefer, 1936, E. festivus Lindberg,
1955, E. leptacanthus Kiefer, 1956, E. alticola Kiefer,
1957, E. neumani titicacae Kiefer, 1957, E. demacedoi
Lindberg, 1957, E. serrulatus montanus Harris, 1978, E.
subciliatus Dussart, 1984, E. pseudoensifer Dussart, 1984,
E. conrowae Reid, 1992, E. torresphilipi Suárez-Morales,
2004, E. chihuahuensis Suárez-Morales and Walsh, 2009,
and E. elegans.
Methods
Panbiogeography is an approach originally proposed by
Croizat (1958, 1964), that aims to analyze the spatial and
temporal distribution patterns of organisms based on a
correlation between the history of the biota and the history of
the Earth. The panbiogeographic approach is based on three
basic concepts: 1) the individual track, which represents the
spatial coordinates of the taxon in space (the geographical
area in which its evolution has taken place), operationally
corresponding to a line that connects localities where a
species or supraspecific taxon is distributed; after these
tracks are constructed, their orientation or direction can
be determined using a baseline (geological feature such as
an ocean or sea basin, or other major tectonic structure,
crossed by the track), main massing (a concentration of
MERCADO-SALAS ET AL.: DISTRIBUTION OF THE AMERICAN EUCYCLOPS
numerical, genetical or morphological diversity within a
taxon in a given area), or phylogenetic evidence available
(directing the track from the most primitive to the most
derivated taxa); 2) the generalized track, which is the
distribution pattern obtained from the overlapping of at least
two individual tracks, indicates the existence of a shared
biogeographic history of the biota and the areas involved;
and 3) the node, which is a complex area where two or more
generalized tracks superimpose and are usually interpreted
as a tectonic/biotic convergence area (Morrone, 2009). For
details of the panbiogeographic methodology see Morrone
and Crisci (1995) and Morrone (2009).
To perform the panbiogeographic analysis, individual
databases were made for each species. Individual tracks
were obtained by using ArcView GIS 3.2 software and the
extension Trazos 2004 (Rojas, 2004); generalized tracks
were obtained superimposing the individual tracks. The
biogeographical system used in this work follows Morrone
(2006).
R ESULTS
We obtained 19 individual tracks (Figs. 1 and 2). Ten
species are restricted to South America, one is exclusive
to Cuba, and seven species are restricted to North America. Mexico is the country with most species recorded
(12), followed by Colombia (7), Brazil, the United States,
Venezuela, and Ecuador (6), Argentina and Peru (4), Chile,
Canada, Nicaragua, and Paraguay (3), Uruguay, Bolivia, and
Cuba (2), and Haiti, Jamaica, Costa Rica, Honduras, and
French Guiana (1). Considering the biogeographic regions
and transitional areas recognized for the Americas (Morrone, 2006), the Neotropical region shows the highest number of species recorded (18), with E. demacedoi, E. neotropicus, E. siloii, E. neumani neumani, and E. silvestri being
endemic to it. The Nearctic region is the second in species
richness with 14 species, with E. macrurus, E. cuatrocienegas, E. borealis, and E. agilis monticola being Nearctic endemics. The third richest area is the Mexican Transition
Zone, with 10 species, none of them endemic. The South
American Transition Zone has seven species, E. breviramatus being the only endemic. Finally, the Andean region
includes only two species (E. pectinifer and E. ensifer),
widespread in other regions as well. Eucyclops pectinifer is
the most widely distributed species, found in all the regions
and transition zones.
Based on the superposition of the individual tracks, two
generalized tracks were found (Fig. 2):
1) Western Amazonian track: southern Peru, eastern
Brazil, and central Colombia. It corresponds to the Amazonian subregion (Pantanal, Madeira, Napo, Imeri, and
Venezuelan Llanos biogeographic provinces) and the South
American Transition Zone (Puna province). Species and
subspecies supporting this track are E. alticola, E. demacedoi, and E. neumani titicacae.
2) Mesoamerican-Northwestern South American track:
central Colombia, Central America, and northeastern Mexico. It corresponds to the Neotropical region, the Mexican
Transition Zone, and the Nearctic region, in the Venezuelan Llanos, North Andean Paramo, Magdalena, Cauca,
Choco, Eastern Central America, Western Panamanian,
459
Mexican Pacific Coast, Chiapas, Sierra Madre del Sur, Balsas Basin, Transmexican Volcanic Belt, Sierra Madre Occidental, and Mexican Plateau biogeographic provinces. The
species supporting this track are E. bondi, E. chihuahuensis,
E. delachauxi, E. festivus, E. leptacanthus, E. pseudoensifer,
and E. torresphilipi.
One node was found in Colombia (Neotropical region), in
the area where both generalized tracks intersect.
D ISCUSSION
The knowledge of the diversity of the American species
of Eucyclops is still growing; several species have been
described recently from different environments. In South
America, many species with a restricted distribution were
described from explorations performed between 1920 and
1990 (Kiefer, 1926, 1929, 1931, 1934, 1936, 1956; Lindberg, 1955; Dussart, 1984; Reid, 1985; Dussart and Frutos, 1986; Defaye and Dussart, 1988). In North America,
the distribution of several species that were previously considered cosmopolitan, e.g., E. serrulatus and E. pseudoensifer, was reevaluated, revealing some new species, probably
with restricted distribution; this suggests that the diversity
of the genus may be underestimated (Reid, 1992; SuárezMorales, 2004; Alekseev et al., 2006; Suárez-Morales and
Walsh, 2009). In Central America, the knowledge about this
genus is still quite limited because surveys in the area are relatively scarce (Suárez-Morales, 2004; Alekseev et al., 2006;
Suárez-Morales and Walsh, 2009).
The distributional patterns of Eucyclops involve two
cenocrons (sensu Morrone, 2009): one Holarctic and the
other Paleotropical. The species of the Holarctic cenocron
have dispersed to the Americas by the Thulean bridge
(connecting North America and Europe) and the Beringian
bridge (connecting North America and eastern Asia) that
existed in the early Eocene when weather conditions were
warmer and wetter (San Martín et al., 2001; Wyngaard et
al., 2009). The individual track of E. pectinifer (Fig. 1)
supports this hypothesis, as the species complex to which
it belongs occurs in Eurasia. According to Alekseev et al.
(2006), E. serrulatus is distributed in North Africa, the
Mediterranean region, Europe, Russia, and probably extends
to Central Asia. The ancestor of E. pectinifer may have
dispersed to North America and then spread southwards.
There is a disjunction between North and South America,
as there are no records of this species in Central America,
linking the individual track from southern Mexico (near the
borderline with Guatemala and Belize) with the Galapagos
Islands. A similar connection through the Galapagos has
described for other organisms such as the staphylinid beetle
genus Rothium (Grehan, 2001). Croizat (1958) considered
the Galapagos Islands as a node that includes the intersection
of three tracks. He predicted that the line of the west
coast of America to the Galapagos previously included
and extended to Chile and Hawaii. Pacific tracks identified
by Croizat constitute evidence of geological connections
with East Asia, which resulted in a composed origin of
North and South America, though the union of lands in the
Pacific, Gondwana, and Laurasia. Grehan (2001), based on
panbiogeographic and geological evidence, considered that
at least some elements of the Galapagos biota have derived
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Fig. 1. Individual panbiogeographic tracks of American species of Eucyclops. Eucyclops pectinifer, E. elegans, E. bondi, E. leptacanthus, E. prionophorus,
E. pseudoensifer.
from the Pacific Islands biota that where in contact with the
Galapagos Islands, and then moved eastward to finally crash
with North or South America.
Ancestors of the South American Eucyclops evolved in
the Afro-Brazilian tropical fragment of Gondwana and diverged following its breakup in the Cretaceous. This has
been proposed by Banarescu (1992), Suárez-Morales et al.
(2005), Hołyńska (2006), and Wyngaard et al. (2009) for
other copepods, such as calanoids (Boeckellidae and Diaptomidae, considered to represent ancient freshwater lineages)
and the cyclopoid genus Mesocyclops. Most of the species
of Eucyclops are distributed in Africa, Central and Southern Asia, Australia, South America, and Mexico, supporting
the hypothesis that the origin of the genus is in the AfroBrazilian tropical fragment of Gondwana. This hypothesis
is also supported if we consider the main massing concept
(center of great diversity), because the distribution of most
species of Eucyclops in the Americas extends from northern-
central Mexico through Colombia, representing 42% of the
species (see individual tracks of E. bondi, E. leptacanthus, E.
pseudoensifer, E. torresphilipi, E. prionophorus, E. ensifer,
E. conrowae, and E. serrulatus montanus), and five species
have north-south tracks within South America (E. alticola,
E. neumani titicacae, E. demacedoi, E. silvestri, and E. neumani neumani).
The proximity of the individual tracks of E. neumani neumani, E. subciliatus, E. elegans, E. pectinifer, and E. silvestri
in the vicinity of northeastern Argentina and southern Brazil
(Chaco, Pampa, and Parana Forest biogeographic provinces)
can be related to the separation of the Paraguay and Paraná
rivers from the Amazonas basin. Castellanos (1965) postulated that the Paraguay River was a former tributary of the
Amazon River, and then splitted when the Andean orogeny
created an area of fracture where the Paraguay and Parana
rivers currently flow, thus changing its drainage to the La
Plata Basin, and acting as a vicariant event separating the
MERCADO-SALAS ET AL.: DISTRIBUTION OF THE AMERICAN EUCYCLOPS
461
Fig. 2. A-C, Individual panbiogeographic tracks of American species of Eucyclops. Eucyclops conrowae, E. torresphilipi, E. neumani titicacae, E.
demacedoi, E. ensifer, E. neumani neumani, E. alticola, E. festivus, E. serrulatus montanus, E. silvestri, E. subciliatus, E. chihuahuensis. Generalized
tracks: 1) Western Amazonian; 2) Mesoamerican-Northwestern South American. The circle with the “x” represents the node.
Amazon and La Plata biotas. The Chaco and Pampa biogeographic provinces belong to the Chacoan subregion, which
includes the Caattinga Province, where the oldest copepod
fossil was found (Kabatarina pattersoni, a parasite of a
teleost fish from the late Cretaceous about 69 m.y.a.) (Huys
and Boxshall, 1991; Lange and Schram, 1999).
The Western Amazonian track is defined by the exclusive presence of three species, E. alticola, E. demacedoi,
and E. neumani titicacae. It is localized mainly in the Amazonian subregion, the largest of the Neotropical subregions.
The history of the Amazonian biota has been reconstructed
differently by different authors. One of the first explanations
was provided by Wallace (1852), who considered that the
rivers of the Amazonian basin acted as barriers. Contrariwise, Arzamendia and Giraudo (2009) recently considered
the rivers as dispersal corridors for snake species. Antonelli
et al. (2009) suggested that the uplift of the tropical Andes in the Neogene strongly affected the history of South
America, changing the course of the Amazon River from a
northwestwards flow to the modern pattern flowing to the
Atlantic. This event changed the climate of the region by
forming the only barrier to atmospheric circulation in the
Southern hemisphere. Wesseling (2006) proposed another
explanation, who based his theory on the Miocene “Pebas
lake/wetland system,” viz., a shallow system of lakes and
wetlands that straddled the Equator in western Amazonia between 9 and 23 m.y.a. This system resulted from the uplift
of the eastern Cordillera in the Central Andes that caused
the western Amazonia to become flooded. The inland sea
thus formed acted as a barrier between the Andes and lowland Amazonia. Aquatic conditions seem to have persisted in
western-central Amazonia until 7 m.y.a., when the modern
Amazon system came into existence; this event could represent the vicariant event associated with the Western Amazonian track.
If divided into two sectors, the Mesoamerican-Northwestern South American generalized track can be compared to that found for other taxa. The northern part in-
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cludes mainly the Mexican Plateau and part of the Sierra
Madre Occidental biogeographical provinces, and agrees
with the Mexican Mountain general track proposed by Abrahamovich et al. (2004) for hymenopteran insects and the
western part of the northern Mexican generalized track proposed by Rosas-Valdéz and Pérez-Ponce de León (2008)
for helminth parasites of ictalurid fishes. It covers almost
all arid areas of north-central Mexico, formed between the
Late Oligocene and Middle Miocene, and was part of a
general trend toward greater aridity resulting from climate
change associated to intense volcanic activity and tectonics
of the Cenozoic, when the Rocky Mountains, the Mexican
and Central-American Plateaus, and the Sierras Madre were
formed. The formation of the Sierra Madre Occidental and
Sierra Madre Oriental during the Eocene and up until the
middle Miocene provided a new barrier to the atmospheric
flow, blocking the masses of warm, moist air from the Pacific Ocean and the Gulf of Mexico, and causing a severe
drought and aridity in the Mexican Plateau. The Miocene
climate change segregated the species along latitudinal and
longitudinal gradients, thus favoring radiation processes in
some lineages (Devitt, 2006). It has been suggested that
some areas (such as Cuatro Cienegas in Coahuila) functioned as refugia during the Pleistocene glaciations (Banarescu, 1992). The southern part of this portion includes
the intersection between the Balsas Basin, Transmexican
Volcanic Belt, and Sierra Madre del Sur biogeographic
provinces, where nodes were identified for arthropods (Morrone and Márquez, 2008; Yáñez-Ordóñez et al., 2008) and
mammals (Escalante et al., 2004). This pattern agrees with
the Mesoamerican generalized track found by Asiain et al.
(2010) for beetles of the genera Agrodes and Plochionocerus. It also coincides with the Meridional distribution
pattern of Maya-Martínez et al. (2011), based on Charaxinae, the Southern Mesoamerican track found by Abrahamovich et al. (2004) for species of Bombus, the Septentrional Mesoamerican and Meridional Mesoamerican generalized tracks described by Márquez and Morrone (2003) for
the staphylinids Heterolinus and Homalolinus, and also with
the Southern generalized track of Morrone and Márquez
(2001), based on beetles. These authors postulated different vicariant events, which occurred in different periods of
time, including the development of the marine barrier of the
Isthmus of Tehuantepec, the emergence of the mountains in
Chiapas, Guatemala, Honduras, and Nicaragua, the development of the Nicaraguan lowlands, and the highlands of
Costa Rica and Panama, and finally the development and
closure of the marine barrier represented by the Isthmus of
Panama. The intrusion of copepod species into Mexico from
the south has been associated with the Usumacinta basin,
mainly by the development of rivers and terraces in the Pleistocene (Gutiérrez-Aguirre and Suárez-Morales, 2001).
The node found in this study appears to be the result
from the mixture of Nearctic and Neotropical biotas after
the closure of the Panama Isthmus, when the connection of
the two subcontinents was consolidated. As already stated
(Menu-Marque et al., 2000), it is important to emphasize
that biogeographic patterns are the consequence not only of
vicariant events, but dispersal and/or extinction processes.
We attempted herein to clarify the distribution of American freshwater copepods using a panbiogeographic approach and covering a wide geographical area. We recognized some of the biogeographic provinces proposed by
Morrone (2006), but there are still many unexplored regions.
Of course, more detailed local and regional records would allow a better understanding of the history and evolution of the
group. A morphological study of the species of Eucyclops in
Mexico is now being conducted in order to correctly define
the species boundaries and clarify their distributional patterns. This study will serve as a platform to define the taxonomic status of the species of Eucyclops in the Americas
and their distributions, which will be analyzed in subsequent
papers.
ACKNOWLEDGEMENTS
We deeply appreciate the comments and help from Juan J. Schmitter-Soto.
We also appreciate the technical support from Holger Weissenberger, who
kindly helped us with the use of Geographic Information Systems. Rosa
Ma. Hernández kindly provided the records of specimens deposited in the
collection of ECOSUR. This contribution is part of the graduate work
of the senior author and was supported by CONACYT project 133404Investigación Científica Básica 2009.
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R ECEIVED: 14 June 2011.
ACCEPTED: 29 October 2011.